Seasonal and spatial dynamics of Centropages typicus and C. hamatus in the western North Atlantic

Centropages typicus and Centropages hamatus are two of the most abundant copepods on the continental shelf in the mid-Atlantic region of the western North Atlantic. Their range extends from the Scotian Shelf (C. typicus) and the Grand Banks (C. hamatus) in the north to Cape Hatteras in the south. South of Cape Hatteras they have only been observed in inshore waters of North and South Carolina and not offshore on the continental shelf or in coastal waters of eastern Florida. However, C. hamatus has been observed in western Florida. Abundances of both species are greatest in inshore regions in the mid-Atlantic Bight with C. hamatus tending to have a more coastal distribution. In this region seasonal variability is low with high abundances from late fall through mid-summer. In the north year-round presence of both species is confined to inshore areas and offshore banks such as Georges Bank, and Browns Bank, Emerald Bank and Western Bank on the Scotian Shelf. In this northern region there is a pronounced seasonal cycle in abundance with high abundances during late summer and fall. Periods of high reproductive rates are closely linked to blooms of large phytoplankton and food availability rather than temperature appears to be controlling population abundances.     

Reproduction,hatching success,and early naupliar survival in Centropages typicus

The broadcast spawner, Centropages typicus, is a very successful copepod species in many coastal areas of the Mediterranean Sea and North Atlantic Ocean. This review assembles the large amount of information on the reproduction and early life history of C. typicus that has emerged since the 1970s and has made this species one of the best-studied copepods, similar in that regard to species of Acartia and Calanus. Observations on mating behavior and the female gametogenic and oogenic cycles are presented, together with information on seasonal cycles of egg production rates in Mediterranean and Atlantic populations from various regions. These studies indicate a strong latitudinal gradient, with continuous reproduction and the main spawning season occurring earlier (late winter/spring) in warmer waters such as the Mediterranean Sea, compared to northern areas such as the North Sea and in the Kattegat, where C. typicus actively reproduces mainly in late summer and fall with reproduction ceasing altogether in winter in the German Bight. These observations strongly suggest that temperature is the controlling factor for reproductive activity in this species. Egg development times are also temperature dependent but do not vary with latitude, and there is as yet no conclusive evidence that diapause egg production occurs in C. typicus. Laboratory experiments have shown that food quantity and quality both affect fecundity and offspring fitness, but most of these studies have focused on diatom and dinoflagellate diets and non-algal prey have been strongly underrepresented, despite their importance for this omnivorous copepod. Large fluctuations in hatching success and naupliar survival have been reported in field surveys and have subsequently been related to maternal feeding history and food quality or toxicity in laboratory experiments. We identify future lines of research that will help to explain the interannual variability in breeding intensity and recruitment of C. typicus in the field based on better understanding of the species’ reproductive biology.     

The feeding ecology of the copepod Centropages typicus (Kröyer)

Here we report on the current knowledge on the feeding ecology of the planktonic copepod Centropages typicus. We describe the foraging process of C. typicus from the detection of prey to their digestion, considering also the effects of several physical variables on the feeding activity of the species. C. typicus is an omnivorous copepod that feeds on a wide spectrum of prey, from small algae (3–4 μm equivalent spherical diameter, ESD) to yolk-sac fish larvae (3.2–3.6 mm length). It uses both suspensivorous and ambush feeding strategies, depending on the characteristics of the prey. In general, C. typicus exhibits selection for large motile prey, such as ciliates or dinoflagellates, both in nature and laboratory, and this selective pattern is enhanced under moderate intensities of turbulence. Daily rations in the field are somewhat lower than those found in the laboratory, which indicates food limitation. This fact, together with the relatively modest capacity of the species to adapt to fluctuations in food availability may explain the geographical distribution of C. typicus, being restricted to near-shelf waters. In general, it does not seem that C. typicus feeding severely impacts planktonic populations. However, the occasional importance of the species in certain ecosystems is also apparent.     

Comparative seasonal dynamics of Centropages typicus at seven coastal monitoring stations in the North Sea,English Channel and Bay of Biscay

We review current knowledge and understanding of the biology and ecology of Centropages typicus in the European shelf-seas (e.g. North Sea, English Channel and Bay of Biscay). Our study is based on observations at seven coastal time-series stations as well as on the Continuous Plankton Recorder dataset. This paper focuses on the influence of the environmental parameters (e.g. temperature and Chla) on the life cycle and distribution of C. typicus and provides a comparison with its congeneric species C. hamatus and C. chierchiae in the study area. Data on abundance, seasonality and egg production have been used to define the temperature and chlorophyll optima for occurrence and reproduction of Centropages spp. within this region of the European shelf-seas.     

Temporal variability of Centropages typicus in the Mediterranean Sea over seasonal-to-decadal scales

Centropages typicus is one of the most common, abundant and best studied calanoid copepods in neritic waters of the Mediterranean Sea, which means it can provide useful information about the long-term dynamics of the Mediterranean epipelagic ecosystem. This paper presents the first comparative overview of the seasonal and long-term variability of C. typicus in different Mediterranean regions. This review is based on quantitative information from the published literature and novel data from five ongoing zooplankton time-series carried out in the Mallorca Island (Balearic Sea), the Bay of Villefranche (Ligurian Sea), the Gulf of Naples (Tyrrhenian Sea), the Gulf of Trieste (North Adriatic Sea), and the Saronikos Gulf (Aegean Sea). In most Mediterranean regions, C. typicus has a perennial occurrence, with peaks of abundance that reflect the succession of different generations. Throughout the Mediterranean, the annual cycle of C. typicus is characterized by minima in winter and major peaks in April–June, which is earlier than those observed in European Atlantic waters, where the peaks are more frequently recorded in summer and fall. In the regions investigated, the annual cycle shows remarkable similarities in terms of timing, but notable differences in the peak height; populations are far more abundant in coastal north-western regions and less abundant in the eastern basin. In the long-term, changes in C. typicus phenology observed in the Bay of Villefranche and in the Gulf of Naples are related to the North Atlantic Oscillation (NAO) index. In these two regions, the species responds to climate forcing similarly in terms of average seasonal patterns (bi-modal patterns in years of positive NAO, unimodal patterns in years of negative NAO) but oppositely in terms of quantity, indicating different influence of the NAO on the two regions. At decadal scales, C. typicus populations show high interannual variability with marked geographical differences. In some areas, the patterns are clearly characterized by alternate phases of higher and lower annual abundances, at higher frequency (mainly 1–2 years) in the Gulf of Naples, and lower frequency in the Saronikos Gulf (mainly 4–5 years) and in the Gulf of Trieste (mainly 5–6 years). Synchronous phases of increasing or decreasing abundance are discernable only for a few sites and short periods, for example from 1998 to 2000 in the Gulf of Naples, Gulf of Trieste and Saronikos Gulf. The regional differences observed in the long-term patterns of C. typicus populations suggest that the temporal dynamics of this species are significantly more affected by local conditions than by any possible common driving force acting at basin scale through teleconnections.     

The biology and ecology of Centropages typicus: An introduction

We briefly introduce the different chapters of this special issue which review the current knowledge and understanding of the biology and ecology of Centropages typicus obtained over the last 30 years. The papers are grouped into two major theme areas: the first five papers review behaviour, feeding, metabolism, growth and development, and reproduction, and the four following papers have a regional focus dealing with temporal and spatial distribution in the Mediterranean Sea and in the North Atlantic shelf areas. The findings reviewed in the two groups of papers converge to explain why C. typicus is such a very successful copepod species in these coastal areas. In the different chapters, similarities and differences with congeneric species (C. velificatus, C. hamatus, C. brachiatus, C. chierchiae, C. furcatus, and C. abdominalis) have been explored where these are known.     

Development and growth rates of Centropages typicus

This paper presents an updated review of our knowledge about Centropages typicus development and growth based on a synthesis of all data found in the literature on stage durations, development rates, size, weight, and C:N ratios. Developmental and growth characteristics have been recorded during field time series and cruises, as well as in controlled laboratory experiments. A few laboratory studies have contributed to an understanding of the impact of food quantity, food quality and temperature on growth and development.C. typicus has a sigmoidal development pattern when the contribution of the different stages to total development time is considered, CIV and CV having much longer durations than preceding stages. The development times of this species are closely related to temperature with a nearly equiproportional development. In turn, food conditions, both in quantity and quality, have less effect on development than on growth rate, leading to large variations in weight within stage, even larger than those predicted from changes in length and biovolume. Actual measurements of weight and biochemical composition are required to better understand the growth of C. typicus in relation to food composition. As a consequence, our review shows that weight–length relationships obtained in particular environmental conditions may be inadequate for others.The combination of a temperature-dependent development rate and a food-dependent growth rate may explain why shelf regions are a more favourable environment for C. typicus as compared to offshore regions, as they support large females, production of more and better quality eggs, and probably better survival of the offspring.     

东海富营养化封闭海水养殖海域浮游植物分布与控制因子

The distribution of phytoplankton and its correlation with environmental factors were studied monthly during August 2012 to July 2013 in the Yantian Bay. A total of 147 taxa of phytoplankton were identified, and the average abundance was in the range of 0.57×10~4 to 7.73×10~4 cell/L. A total of 19 species dominated the phytoplankton assemblages, and several species that are widely reported to be responsible for microalgae blooms were the absolutely dominant species, such as Skeletonema costatum, Navicula sp., Thalassionema nitzschioides,Pleurosigma sp., and Licmophora abbreviata. The monthly variabilities in phytoplankton abundance could be explained by water temperature, dissolved oxygen, salinity, dissolved inorganic nitrogen(DIN), and suspended solids. The results of a redundancy analysis showed that p H and nutrients, including DIN and silicate(SiO_4), were the most important environmental factors controlling phytoplankton assemblages in specific months. It was found that nutrients and pH levels that were mainly influenced by mariculture played a vital role in influencing the variation of phytoplankton assemblages in the Yantian Bay. Thus, a reduction of mariculture activities would be an effective way to control microalgae blooms in an enclosed and intensively eutrophic bay.     

Metabolism of Centropages species in the Mediterranean Sea and the North Atlantic Ocean

Information on the metabolism rates of Centropages typicus and congeneric species (C. hamatus, C. furcatus, C. brachiatus and C. abdominalis) in neritic areas of the Mediterranean Sea, the North Atlantic Ocean and the Pacific Ocean are reported here. Respiration rates and excretion rates are strongly influenced by abiotic (i.e. temperature, salinity) and biotic factors (i.e. food availability and composition). Differences in the response of respiratory rates to temperature of acclimated, acclimatized and adapted individuals are clearly observed among regions of the Mediterranean Sea and the West and East shores of the Atlantic Ocean. Food supply also strongly affects respiration and excretion rates, as well as the size, sex and stage development of the individuals. The co-measurement of these two rates allows confirmation of the omnivory or carnivory oriented feeding habits of these species. The role of this neritic genus in coastal environment is also discussed.     

Spatio-temporal changes in the distribution of phytopigments and phytoplanktonic groups at the Porcupine Abyssal Plain (PAP) site

We have made a comprehensive study of pigment distributions and microscopically determined phytoplankton abundances within the Porcupine Abyssal Plain (PAP) location in the North Atlantic to better understand phytoplankton variability, and make some suggestions regarding the composition of the material falling to the sea bed and its impacts on benthic organisms such as Amperima rosea. The area has been the focus of many studies of ocean fluxes and benthic communities over recent years, but little attention has been given to the spatio-temporal variability in the surface waters. Dawn casts over a 12-day period at the PAP mooring site (48.83°N 16.5°W) revealed the presence of only one species, the diatom Actinocyclus exiguus, at bloom concentrations for just 5 days. Smaller populations of other diatoms and the dinoflagellates Gymnodinium and Gyrodinium were also present at this time. Following this 5-day interval, a mixed population of small–sized dinoflagellates, prymnesiophytes, prasinophytes, chrysophytes and cyanobacteria occurred. It is clear from concomitant CTD/bottle surveys that rapid changes in phytoplankton community structure at a fixed time series position do not necessarily reflect a degradation or manifestation of one particular species but rather represent the movement of eddies and other water masses within very short timescales. These cause substantial variability in the species class and size fraction that may explain the variability in carbon export that has been seen at the PAP site. We also make some suggestions on the variable composition of the material falling to the seabed and its impact on benthic organisms such as Amperima rosea.     

Long-term changes of phytoplankton community in Xiagu waters of Xiamen, China

Long-term changes of phytoplankton community by water sampling method in Xiagu Sea waters of Xiamen, China, were investigated in this study. Species composition of the phytoplankton community in these waters changed greatly since the 1950s. The numbers of Dinophyta species increased significantly, although Bacillariophyta species are generally dominant. The succession of dominant species in phytoplankton community is obvious:large-size dominant species such as Biddulphia sinensis of the 1950s were gradually replaced by small-size ones such as Cyclotella striata and Nitzschia closterium, and species that still maintain dominant such as Skeletonema costatum are also small ones, leading the whole phytoplankton community of smaller size. Cell density of phytoplankton community increased greatly, among which cell density of the most dominant species Skeletonema costatum have been increasing in exponent function. Margalef index of phytoplankton community decreased, indicating decline of biodiversity of the community, and dominant character of Skeletonema costatum increased. Generally, the structure of the entire phytoplankton community is becoming more and more singular and unstable, which makes the occurrence of red tides more frequent. The succession in the phytoplankton community is related to the long-term changes in marine environment, influenced by human activities and global climate changes, especially the increases of nutrient content.     

Environmental correlates of large-scale spatial variation in the δ13C of marine animals

Carbon stable isotopes can be used to trace the sources of energy supporting food chains and to estimate the contribution of different sources to a consumer's diet. However, the δ¹³C signature of a consumer is not sufficient to infer source without an appropriate isotopic baseline, because there is no way to determine if differences in consumer δ¹³C reflect source changes or baseline variation. Describing isotopic baselines is a considerable challenge when applying stable isotope techniques at large spatial scales and/or to interconnected food chains in open marine environments. One approach is to use filter-feeding consumers to integrate the high frequency and small-scale variation in the isotopic signature of phytoplankton and provide a surrogate baseline, but it can be difficult to sample a single consumer species at large spatial scales owing to rarity and/or discontinuous distribution. Here, we use the isotopic signature of a widely distributed filter-feeder (the queen scallop Aequipecten opercularis) in the north-eastern Atlantic to develop a model linking base δ¹³C to environmental variables. Remarkably, a single variable model based on bottom temperature has good predictive power and predicts scallop δ¹³C with mean error of only 0.6‰ (3%). When the model was used to predict an isotopic baseline in parts of the overall study region where scallop were not consistently sampled, the model accounted for 76% and 79% of the large-scale spatial variability (10¹–10⁴ km) of the δ¹³C of two fish species (dab Limanda limanda and whiting Merlangus merlangius) and 44% of the δ¹³C variability in a mixed fish community. The results show that source studies would be significantly biased if a single baseline were applied to food webs at larger scales. Further, when baseline δ¹³C cannot be directly measured, a calculated baseline value can eliminate a large proportion of the unexplained variation in δ¹³C at higher trophic levels.     

Phytoplankton blooms in the Urdaibai estuary during summer: Physico-chemical conditions and taxa involved

In the Urdaibai estuary, despite its small volume in relation to the tidal prism, phytoplankton grows massively in the upper and intermediate zones of the estuary during summer when rainfall is low to moderate. Data obtained in 23 sampling transects along the longitudinal axis of the estuary, undertaken in July 1993 and August 1994, showed three distinct zones in terms of the phytoplankton species composition: in the upper zone the phytoplankton assemblage was dominated by the diatoms Cyclotella spp., the dinoflagellate Glenodinium foliaceum, cryptophytes and euglenophytes; in the intermediate zone the diatom Chaetoceros ceratosporus, the dinoflagellates Peridinium quinquecorne and Prorocentrum minimum and cryptophytes were the most abundant; in the lower zone diatoms such as Leptocylindricus danicus and Skeletonema costatum together with the dinoflagellates Prorocentrum spp. were dominant. The distribution and abundance of these phytoplankton forms is discussed in relation to variations in salinity, water column stratification, nutrients and temperature, which in turn vary mainly as a function of meteorological conditions (i.e. rainfall, irradiance) and tidal amplitude. The longitudinal distribution of phytoplankton cells suggests that the diatoms Cyclotella spp. and Chaetoceros ceratosporus undergo a greater dispersion along the estuary during rain events than the flagellates, thus contributing to the transport of silica to the outer areas of the estuary.     

The coccolithophore summer–autumn assemblage in the upwelling waters of Portugal: Patterns of mesoscale distribution (1985–2005)

A study of the mesoscale distribution of phytoplankton communities observed along the Portuguese upwelling coast in 1985–1986 made it possible to identify an assemblage composed by the coccolithophores Helicosphaera carteri, Syracosphaera pulchra and Coronosphaera mediterranea. The assemblage was consistently present in the coast in the following years (1991, 1992, 1994, 2003 and 2005) and from 1992 onwards, Rhabdosphaera clavigera was recognised as being part of it, although in low concentrations During late summer, this group of species showed higher affinities with upwelling waters of subtropical origin (ENACWst) on the SW coast. At the end of the upwelling season, in autumn, the assemblage was advected to the NW shelf due to the intensification of the poleward surface circulation or meridional seasonal changes of environmental conditions. The species maximum abundances were not coincident in space: H. carteri developed at the central part of the western coast while blooms of S. pulchra and C. mediterranea generally occurred further south, such as at the upwelling centre of cape S. Vicente. Syracosphaera pulchra blooms were characterised by lower concentrations of C. mediterranea and vice-versa. The above patterns were recurrent along the years but the relative abundance of each species appeared related to the interannual variability of physical conditions as the upwelling.     

Abundance,community composition,size distribution,and production rates of tintinnids in Narragansett Bay,Rhode Island

The abundance of tintinnid ciliates in lower Narragansett Bay was measured at weekly intervals over the period 1980–1982. Twenty-nine species representing nine genera ranged in abundance from 10¹ to 10⁵ tintinnids l⁻¹. Tintinnopsis was the most numerous genus in terms of numbers of species and individuals. Total abundance increased with water temperature above 6°C, and with nanoplankton chlorophyll a (<10 μm and <5 μm chl a) averaged over the water column. Exceptions occurred during blooms of phytoplankton previously demonstrated to be poor food for tintinnids (Olisthodiscus, Thalassiosira). Tintinnids aggregated near the bottom during periods of low nanoplankton chl a. Abundances and distributions were not correlated with particulate organic carbon. Seasonal changes in length and oral diameter of loricas of individual species were inversely related to temperature. Oral diameter was a more constant feature of lorica morphology than length. Seasonal patterns in the mean oral diameter of the lorica of all species reflected decreases in the oral diameter of individual species with increasing temperature, and changes from small to large species coincident with similar shifts in the size of phytoplankton. The production rate of tintinnids was 3·3 mgCl⁻¹ year⁻¹, equivalent to the ingestion of 26% of total annual net primary production and 52% of the estimated production of <10 μm phytoplankton.     

A quantitative analysis of sources for summertime phytoplankton variability over 18 years in the South Shetland Islands (Antarctica) region

Eighteen years of summertime hydrographic and chlorophyll-a (Chl-a) data (～2700 stations) from the South Shetland Islands (Antarctica) region show that a “bell-shaped” (unimodal) distribution of phytoplankton biomass results annually when plotted against the inshore to offshore gradient in surface salinity. The maximum for this unimodal Chl-a distribution corresponds with a shallow upper mixed layer (UML) in iron-rich waters that occurs at salinities ～34. Methods of gradient analysis are used to distinguish sources of variability for bloom development among years. The control of phytoplankton biomass is resolved across the salinity gradient that separates the co-limiting conditions of deep UML depths and low-iron concentrations as opposing end-members. Chlorophyll-fluorescence yield data (a proxy for Fe-stress) showed that at salinities ～34, phytoplankton biomass was unlikely to be limited by Fe. Instead, blooming at salinities ～34 (1.3±1 mg Chl-a m^?3) co-varied with shallow UML depths (41±19 m) that occurred as a function of higher UML temperature (1.5±0.5 °C) among years, and is evidence that atmospheric climate variability impacts summertime phytoplankton biomass and production in this Southern Ocean seascape.     

2010年西北冰洋浮游植物区域差异与环境关系

Global warming has caused Arctic sea ice to rapidly retreat,which is affecting phytoplankton,the primary producers at the base of the food chain,as well as the entire ecosystem.However,few studies with large spatial scales related to the Arctic Basin at high latitude have been conducted.This study aimed to investigate the relationship between changes in phytoplankton community structure and ice conditions.Fifty surface and 41 vertically stratified water samples from the western Arctic Ocean(67.0°–88°26′N,152°–178°54′W) were collected by the Chinese icebreaker R/V Xuelong from July 20 to August 30,2010 during China's fourth Arctic expedition.Using these samples,the species composition,spatial distribution,and regional disparities of phytoplankton during different stages of ice melt were assessed.A total of 157 phytoplankton taxa(5 μm) belonging to 69 genera were identified in the study area.The most abundant species were Navicula pelagica and Thalassiosira nordenskioeldii,accounting for 31.23% and 14.12% of the total phytoplankton abundance,respectively.The average abundance during the departure trip and the return trip were 797.07×10~2 cells/L and 84.94×10~2 cells/L,respectively.The highest abundance was observed at Sta.R09 in the north of Herald Shoal,where Navicula pelagica was the dominant species accounting for 59.42% of the abundance.The vertical distribution of phytoplankton abundance displayed regional differences,and the maximum abundances were confined to the lower layers of the euphotic zone near the layers of the halocline,thermocline,and nutricline.The species abundance of phytoplankton decreased from the low-latitude shelf to the high-latitude basin on both the departure and return trips.The phytoplankton community structure in the shallow continental shelf changed markedly during different stages of ice melt,and there was shift in dominant species from centric to pennate diatoms.Results of canonical correspondence analysis(CCA) showed that there were two distinct communities of phytoplankton in the western Arctic Ocean,and water temperature,ice coverage and silicate concentration were the most important environmental factors affecting phytoplankton distribution in the surveyed sea.These findings will help predict the responses of phytoplankton to the rapid melting of Arctic sea ice.     

2018年烟台四十里湾及其邻近海域网采浮游植物群落结构的时空变化

本文基于2018年5月（春季）、8月（夏季）、11月（秋季）和2019年1月（冬季）季度航次调查,采用冗余分析（RDA）和Bray-Curtis相似性聚类分析方法,讨论了烟台四十里湾及其邻近海域网采浮游植物丰度、优势种组成、多样性指数、均匀度指数的分布特征以及浮游植物与环境因子的关系。本年度调查共发现浮游植物4门69属116种,其中硅藻门50属85种,甲藻门15属27种,着色鞭毛藻门3属3种,未定类1种（三深碎裂藻,Ebria tripartita）。浮游植物丰度在8月份达到年度最高值,为2.89×10⁸cells/m³,秋季次之,春季最低,春、秋、冬季浮游植物丰度均为~10⁵cells/m³。硅藻门在4个季节中均占绝对优势,除春季出现甲藻门（夜光藻）优势种外,其余季节浮游植物优势种均为硅藻门。相邻季节浮游植物优势种更替明显,未出现4个季节共有的优势种。浮游植物多样性和均匀度指数均在夏季达到最低值,分别为1.58和0.31。RDA排序分析显示,不同季节影响浮游植物群落分布的环境因子不同,总体来讲,温度、溶解氧浓度、SiO₃^2-浓度和NO₃^-浓度对浮游植物的丰度和平面分布有显著影响。Bray-Curtis相似性聚类结果显示,2018年每个季节浮游植物群落分布均可分为两个类群,其分布受营养盐分布的影响较为明显。     

Variability of Particulate Organic Phosphorus in the Northwestern Part of the Black Sea

Based on long-term (1985–1995) monitoring data, the paper considers the peculiarities of seasonal variability in the spatial and vertical distribution of particulate organic phosphorus (Р_POM) in the surface layer and in the photosynthetic zone in the northwestern Black Sea. Regression equations, experimental data, and satellite observations for the chlorophyll a concentration allowed us to evaluate the seasonal longterm (1979–1995) variability in Р_POM in the surface layer and photosynthesis zone. The ratios of the concentrations of particulate organic carbon, nitrogen, phosphorus, and chlorophyll a are calculated and statistical estimates of seasonal changes in the Р_POM in the areas with different degrees of influence of river runoff and water of open seas are obtained. The consistency of intra-annual changes in the concentrations of Р_POM, chlorophyll a, and phytoplankton biomass is shown, which indicates the role of phytoplankton in the formation of Р_POM and in its intra- and interannual variability in the northwestern part of the sea. It is shown that long-term seasonal variations in Р_POM and related changes in the concentration of chlorophyll a depend on the variability of bulk river runoff, the extent of its abundance in the northwestern shelf, and regional hydrometeorological conditions.